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(J. Adams, 1800) (Gastropoda, Prosobranchia) from the Dutch Wadden Sea

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(J. Adams, 1800) (Gastropoda, Prosobranchia) from the Dutch Wadden Sea BASTERIA, Vol. 61: 89-98, 1998 Rissoa membranacea (J. Adams, 1800) (Gastropoda, Prosobranchia) from the Dutch Wadden Sea Gerhard+C. Cadée Netherlands Institute for Sea Research, P.O. Box 59, 1790 AB Den Burg, Texel, The Netherlands Rissoa membranacea disappeared from the Dutch Wadden Sea together with the large subtidal stands L. ofeelgrass Zostera marina in the early 1930s, due to a still not completely understood ‘wasting disease’ in Zostera. A small population of R. membranacea survived together with Zostera marina in inland Bol’ 1981. shells of R. an brackish water ‘de on Texel until Empty membranacea can still be found at these localities. The relatively large size of the top-whorl ofWadden Sea R. membranacea indicates that the larvae In this were lecithotrophic. respect they arecomparable to R. membranacea type B from the Roskilde Fjord of Rehfeldt (1968) and Warén (1996) and R. membranacea to s.s. as described by Verduin (1982b). It is, however, questionablewhether R. membranacea Type A ofRehfeldt and R. labiosa sensu Verduin can be separated from R. membranacea s.s. They differ only in the smaller size of the embryonic whorl indicating planktothrophic larvae. This well indicate different oflarval pelagic might as two types development,lecithotrophic andplanktotrophic in one and the same species R. membranacea s.l. Apical dimensions in samples from the Ria de Arosa (Spain) and from Dutch Eemian deposits show a gradual change from small ‘R. labiosa’ to larger ‘R. membranacea’ values, which supports the merging ofboth nominal species and corroborates Warén’s suggestion of a continuum in larval development in R. membranacea between lecithotrophyand planktotrophy as well as newer ideas of a gradually and easily crossed transition between lecithotrophy and planktotrophy of McEdward (1997). Key words: Gastropoda, Prosobranchia. Rissoidae, Rissoa membranacea, larval development, lecithotrophy, planktotrophy, Wadden Sea, Netherlands. INTRODUCTION Rissoa membranacea J. Adams, 1800) is a small prosobranch gastropod, occurring from Norway and the Baltic near Rugen to the Canary Islands, and probably also in the Mediterranean. It lives associated with marina L.. It is normally eelgrass Zostera an annual species which breeds in summer in the northern part of its distribution (Fretter & Graham, 1978). Egg capsules are deposited preferably on Zostera leaves (Van Goor, 1919; Lebour, 1934; Smidt, 1938). After hatching they grow rapidly. Van Goor (1919) observed 1 mm shells in July and in September they had reached already their max- imum size of 5.5 - 9 mm. Smidt (1938) found an equally rapid growth in the Sound from 1 3.5 - 4 their maximum size for this in mm early August to mm, locality, early fall September. During winter Zostera leaves off and the greater part ofthe R. membrana- small attains the in the cea population perishes, only a part breeding stage following former summer (Van Goor, 1919; Smidt, 1938). In the Zuiderzee R. membranacea was Benthem The very common (Van Goor, 1919;Van Jutting, 1933). species was badly affected by the Zostera disease in NW. Europe in the 1930s. Waren (1996) presents an of its and up-to-date account taxonomy ecology. 90 BASTERIA, Vol. 61, No. 4-6, 1998 The variable shell ofR. membranaceahas induced highly morphology long ago already Schwartz authors to a division in differentspecies and varieties (e.g. von Mohrenstern, 1864). Verduin (1982a,b) meticulously measured numerous Recent shells of R. mem- from Black Sea and the Mediterranean branacea s.l. from European seas ranging the to northern Europe and the Baltic. He was able to distinguish two variable 'species', differed in the size of the whorl. The forms with the smaller which only apical apex he identified R. labiosa those with R. membranacea as (Montagu, 1803), a larger apex as Verduin mentioned Rehfeldt who also had found in R. s.s. (1982b) (1968), two types membranacea. She able relate these difference in larval was to types to a development: with slender and larvae in B with pelagic larvae in type A the apex, non-pelagic type the Rehfeldt's results illustrated the of Thorson larger apex. nicely 'apex theory' (1950), which states that in general a clumsy large apex in gastropods points to non- pelagic development and lecithotrophic larvae (feeding on yolk and not requiring planktonic food), while a narrowly twisted apex points to a pelagic development and larvae. Since Thorson's seminal facul- planktotrophic papers (Thorson, 1946, 1950) tative planktonic feeding has been observed in lecithotrophic larvae e.g. in the nudibranch Phestilla sibogae Bergh, 1905 by Kempf & Hadfield (1985) and Miller (1993) indicating a gradual transition between lecithotrophy and planktotrophy. McEdward and reversible transition between both (1997) suggests a gradual easily feeding types. Also so-called adelphophagy (Bouchet, 1989), the ingestion by developing larvae of nurse eggs or embryo's in the same egg-capsule, has been observed and complicates the picture. Waren (1996) studied > 1,000,000 R. membranacea specimens from all parts of the distributional in and confirms the existence of Rehfeldt's A area western Europe type and B, giving excellent SEM pictures of the difference in the top-whorl. Waren found Rehfeldt's A = Verduin's R. labiosa from northern type (with planktotrophic larvae) Europe to southern Portugal (and presumably in the Mediterranean), whereas Reh- feldt's B = Verduin's R. membranacea in the type s.s. (with development egg capsule, veligers feeding on nurse eggs) is known only from Roskilde fjord, south-west France and the United But he also that the situation be Kingdom. reports may more com- plicated: planktotrophic larvae have been found to consume undeveloped eggs before which enables them size in the and hatching, to grow to a larger egg-capsule probably abbreviate their pelagic period. Waren suggests a continuum between lecithotrophy conditions. and planktotrophy as an adaptation to local Thorson (1950) observed a clear N-S trend in larval development throughout the North Adantic, later named 'Thorson's rule'. He observed only prosobranchs with non- pelagic larvae in East Greenland, to the majority (68%) with pelagic and less (32%) Islands. studies since with non-pelagic larvae at the Canary Numerous 1950 support a general increase in non-pelagic, brooded development with increasing latitude, al- though exceptions have been found in the Antarctic and deep-sea invertebrates (see Levin & for Verduin did not Rehfeldt's Bridge, 1995, a review). (1982b), integrate results in his did he mention how his data fit in with the paper, nor nicely apex theory of Thorson: Verduin observed R. labiosa with the slender and therefore most apex probably with pelagic planktotrophic larvae, from the Mediterranean to Denmark. His R. membranacea with the and larvae did large apex probably lecithotrophic occur more to the north, in the Baltic, the North Sea, the Irish Sea and the Channel, but not south of Bretagne. There is an area of overlap of the two 'species' in which among others The Netherlands but Verduin did material from are situated, unfortunately not study there. This leaves the possibility that both his R. membranacea and R. labiosa once lived Cadee: Rissoa membranaceafrom the Dutch Wadden Sea 91 in The De Boer De Netherlands, as suggested e.g. by & Bruyne (1991) and De Bruyne (1994). R. membranacea s.l. lives in the Dutch no longer Wadden Sea (De Boer & De Bruyne, 1991) and is also extinct in the German and Danish Wadden Sea (Von Nordheim et al., 1996). With the disappearance of large sublittoral stands of Zostera marina in the early 1930s (Polderman & Den Hartog, 1975), R. membranacea and another small gastropod Lacuna vincta living on eelgrass, (Montagu, 1803) (synonym L. divaricata (Fabricius, 1780)) disappeared as well. Van Benthem still mentioned both Jutting (1933) species as very abundant in eelgrass fields, which fauna was well studied by van Goor (1919), but the species had probably already disappeared at the time of publication in 1933. On the other its in Wadden Sea hand, occurrence the was not discovered before the beginning of this century. The first specimens found of R. membranacea date from 1909, collected W.C.Van Heurn Texel and from by on (Vernhout, 1912) 1912 on Griend, collected Van der Sleen In the Wadden Sea shells of both by (1913). now only empty species be found and in small numbers observations and R. can only (personal pers. comm. Also, the small of R. membranacea, which survived Dekker, NIOZ). population among marina in brackish inland 'de Bol' Zostera a water on Texel, has disappeared. 'De Bol' had a connection with the Wadden Sea via an ebb-sluice which opened automatically low tide because then the of the the landside became during pressure water on higher than that from the Wadden Sea water. R. membranacea was discovered here in the 1950s (Swennen, 1955) and repeatedly mentioned and studied since (Reydon & Visser, 1967; Visser, 1968ab; Verhoeven & Van Vierssen, 1977, 1978). After removal of the ebb- sluice during dike reconstructions in 1979, the last Dutchpopulation of Rissoa membranaceabecame extinct (De Jong & De Kroon, 1982; Van der Goes et al. 1996; personal observations). In this I from note report on R. membranacea the Wadden Sea. Some hundred spec- imens collected shells from drift the Wadden Sea south of were as empty along coast the tidal marshes 'De in 1997. Also of shells Schorren', Texel, June a sample empty from the inland brackish water 'de Bol', Texel, sieved from a bottom sample taken in July 1997, was studied. In addition, samples of R. membranacea collected alive in the Wadden Sea between 1909 and 1929 (collection Zoological Museum Amsterdam, ZMA) and from 'de Bol' collected in 1966 and 1968 (kindly provided by G.J.M. Visser, Terschelling, now donated to ZMA) were studied. For comparison Recent specimens from the province of Zeeland and from the Ria de Arosa (Spain) and fossil material from Eemian deposits near Amsterdam were used (see table 1 for data on all samples used).
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